J. Nannoplankton Res. 29 (1), 2007, pp.39-65 © 2007 International Nannoplankton Association
ISSN 1210-8049 Printed by Cambridge University Press, UK

39

New and rarely reported calcareous nannofossils from theLate Cretaceous of coastal Tanzania: outcrop samples andTanzania Drilling Project Sites 5, 9 and 15
Jackie A. Lees
Department of Earth Sciences, University College London, Gower Street, London, WC1E 6BT, UK; j.lees@ucl.ac.uk
Manuscript received 20th February, 2007; revised manuscript accepted 8th March, 2007

Abstract Field sampling and subsurface drilling in Tanzania, in and near to the coastal towns of Lindi andKilwa, and connected to the NERC-funded Tanzania Drilling Project, have provided a wealth of Cretaceous material for analysis. The clay-rich lithologies contain nannofloras that are generally very well preserved and diverse.
Consequently, several new calcareous nannofossil taxa have been discovered, and a number of taxa that are rarely
reported have been recorded. These taxa are described and/or illustrated here. They include: seven new genera(Bilapillus, Duocameratus, Nicholasia, Pearsonia, Singanoa, Tanzanella, Thecatus); 17 new species of heterococcolith (Calciosolenia? huberi, Corollithion karegae, Eiffellithus lindiensis, Gartnerago? coxalliae, Miravetesina
bergenii, Percivalia? dunkleyjonesii, P.? pearsonii, Placozygus banneri, Prediscosphaera mgayae, Rhagodiscus
pancostii, Rotelapillus msakyae, Staurolithites halfanii, S. handleyi, S. ngurumahambaensis, Truncatoscaphus
macmillanii, Zeugrhabdotus blowii, Z. simplex); 22 new species of holococcolith (Bifidalithus mchanae, Bilapillus
wadeae, Calculites cenomanicus, C. cyclops, C. juliae, C. maghredaswampensis, C. paulus, C. proscissus, C. rosalyniae, C. turonicus, Duocameratus leariae, D. sianiae, Lucianorhabdus? boudagherfadeliae, L.? tabernus, L. turris, Metadoga ampulla, Munarinus mkeremei, Nicholasia baileyi, Orastrum robinsonii, Pearsonia ecclesiata,
Tanzanella bownii, Thecatus varolii); four new species of nannolith (Ceratolithoides dongenii, C. ohalloraniae,
Micrantholithus? nicholasii, Singanoa scapus); two new combinations (Cribrosphaerella circula (Risatti, 1973),
Lucianorhabdus compactus (Verbeek, 1976)); and one taxon of elevated status (Ceratolithina capitanea (Burnett,
1997a)).

Keywords Calcareous nannofossils, holococcoliths, Cenomanian, Turonian, Campanian, Maastrichtian, taxonomy, Tanzania
1. Introduction
The Tanzania Drilling Project (TDP) was funded after
three years of fieldwork (1998-2000) in the coastal
Tanzania region, focussed on the towns of Kilwa andLindi (Figure 1). The main aims of the project were to
recover Paleogene and Cretaceous planktonic
foraminifera of unprecedented preservation, as well asorganic biomarkers, from the sediments, primarily inorder to gain meaningful palaeotemperatures and atmospheric CO2 estimates via isotopic analyses (e.g. Pearson etal., 2001; van Dongen et al., 2006). Among the ~500 outcrop samples initially collected were several of
Cretaceous age. To date, those examined for calcareous
nannofossils provide a discontinuous record of the
Albian-Cenomanian, Turonian and CampanianMaastrichtian (Lees, unpubl. data). Furthermore, three ofthe 20 sites drilled (with the collaboration of the Tanzania
Petroleum Development Corporation – TPDC), between
2002 and 2005, also provided Cretaceous material. TDP
Sites 5 and 9 (described in Pearson et al., 2004, 2006;
Nicholas et al., 2006) recovered sediments from aroundthe Campanian/Maastrichtian boundary, whilst TDP15(Nicholas et al., 2006; Pearson et al., in prep.) was drilled
specifically to try to recover the Cenomanian/Turonianboundary, although this proved to be elusive (Lees,
unpubl. data).
Bown (2005), Bown & Dunkley Jones (2006) and

Bown et al. (2007) have published extensively on taxonomic aspects of the Paleogene nannofloras, the dozens of
new species described testifying to the fact that theirs is
exceptionally well preserved material. Preservation of theCretaceous material, especially from the cores, is variable, but overall seems not to be as pristine as that of the
Paleogene cores, basedon both planktonic
foraminiferal and nannofossil scanning electron microscope observations (B.T. Huber,pers. comm., 2006; JAL
pers. obs., 2004-2006).
The calcareous nannofossil preservation in the
Cretaceous is governedby the domination ofclay in the sediment, but
possibly tempered byhaving suffered fromtropicalweathering,even at depth. Having
said that, there are many
new calcareous nanno- Figure 1: General location of thestudy area, coastal Tanzania (after
fossil taxa in the Pearson et al., 2004; Bown, 2005).
Cretaceous material, and Detailed maps can be found in
Nicholas et al. (2006)

Lees

40

this is a preliminary description of the exceptional diversity that I have observed so far.
2. Material
The Upper Cretaceous sediments belong to the
Nangurukuru Formation of the Kilwa Group (as describedin Nicholas et al., 2006). They represent the onset ofincreased subsidence across the shelf, but the sedimentactually varies very little from the Cretaceous throughPaleogene, being predominantly represented by dark clay,
claystones and siltstones (Nicholas et al., 2006). Outcrop
material included herein was collected as spot-samples
from around Kilwa and Lindi (as described below; gridreferences refer to maps shown in Nicholas et al., 2006).
Locality and age details for the outcrop spot-samples
mentioned here are given in Table 1. The nannofossil biozonation of Burnett (1998a) has been applied to thesesamples. (Full biostratigraphical information will be provided in a separate publication.)sampleage

biozoneAround Lindi town, ~UTM 37L 573393, 8894643

Lindi-99-2
Lindi-99-3
Lindi-99-9a
Lindi-99-11
Lindi-99-12

Campanian/Maastrichtian
Early Maastrichtian
Early Maastrichtian
Early Maastrichtian
Early Maastrichtian

UC16
UC17
UC17
UC17
UC18

Nguru-mahamba, SW of Lindi town, UTM 37L 561739, 8891339

Nguru-99-1
Nguru-99-2
Nguru-99-3
Nguru-99-4
Nguru-99-5
Nguru-99-9
Nguru-99-10
Nguru-99-12

Middle-Late Cenomanian
Early Cenomanian
Early Cenomanian
Early CenomanianMiddle-Late Cenomanian
Early Maastrichtian
Late Campanian
Late Maastrichtian

UC3a-b
UC1a-2aUC1a-2aUC1a-2aUC3a-b
UC17
UC15eTP
UC20aTP

Matandu Bridge, west of Kilwa Kivinje (UTM 37L 531458, 9033532)

PP04-K1
PP04-K9
PP04-K17
PP04-K19
PP04-K20

Middle-Late Turonian
Early Maastrichtian
Late Campanian
Late Campanian
Late Campanian

UC9a
UC17
UC15dTPUC15eTP
UC15dTP

Table 1: Outcrop sample locations and biostratigraphy

TDP5 was drilled in 2002 at Machole, next to the roadbetween Lindi and Mtwara, south of Kitulo Hill
(10˚01.646’S, 39˚41.375’E: Pearson et al., 2004). TDP9
was drilled in 2003 near the road junction at
Nangurukuru, WSW of Singino Hill, near Kilwa Kivinje(UTM 37L 538987, 9027049: Pearson et al., 2006).
TDP15 was drilled in 2005 in Maghreda Swamp, NW of
Kilwa Kivinje (UTM 37L 528437, 9046105: Nicholas etal., 2006; Pearson et al., in prep.). Sample notation fromthese cores (e.g. TDP9/29-1, 99cm) refers to the site
(TDP9), followed by the core and section number (29-1)
and then the centimetre depth from the top of that section.

3. Methods
The species described here are based on light-microscope(LM) observation only. This was performed on smearslides, prepared as described in Bown & Young (1998).
Examination was performed on a Zeiss Axio Imager.A1 at
1250x magnification. Digital images were captured with a
Leica DFC280 digital camera at 640x480 pixels, using
Leica FireCam software. Plates were made using Canvas
software on a Mac PowerBook, as follows. In a new
Canvas file, the TIFF images were first acquired (Image>Acquire->TIFF), then scaled down to 25% (Object>Scale) and cropped (Image->Area->Crop), generally to
3x3cm or 2x2cm. The resolution can also be boosted(Image->Area->Resolution->Best). These images werethen arranged into plates, and scale-bars, figure numbers,
taxon names, sample numbers and image-identifying
numbers added. All images on the plates were taken incross-polarised light (XPL), except for those taken withphase-contrast (PC) or bright-field (BF), these being indicated on the plates. Typically, each specimen is figured at
0˚ and ~45˚ to the polarisers.
4. Gross assemblage composition
Species richness is typically very high in this material,
comparing favourably with the global diversity estimatesof Bown et al. (2004), and preservation varies from goodto moderate. Holococcoliths and very small (<3µm) taxa
are frequent components of the assemblages, although
individual holococcolith species are usually rare to few inabundance.
Coccospheres are also frequently represented, particularly those of taxa with placolith morphology (e.g.
Biscutum, which shows a plethora of coccosphere shapes(Pl.5, figs 2-10, 12-15), Cribrosphaerella andDiscorhabdus). Whilst the nannofloras are not particularly dominated by any one taxon, Corollithion,
Cribrosphaerella, Discorhabdus, Prediscosphaera,
Watznaueria and Zeugrhabdotus are the most common
assemblage components. Notably, the typically common
Campanian lucianorhabdids and Calculites obscurus seenelsewhere are absent from, or very rare in, these assemblages. This may indicate that they are not tropical taxa.
The seldom reported Prolatipatella multicarinata
occurs as a rare, yet quite consistent component, in sediments from TDP9 (Upper Campanian-Lower
Maastrichtian). Lees (2002) indicated its status as a
Tropical taxon, and this is supported by its unusually consistent presence in Tanzania. Repagulum parvidentatum isa rare, but sometimes consistent, component of these lowlatitude nannofloras also. This underlines the fact that,whilst an increase in its abundance may be used as aproxy for cooler water, its mere presence in an assemblage may not. Its occurrence in these well-preserved
samples may also involve a preservational bias.
5. Systematic palaeontology
The terminology used in the descriptions below follows

New Late Cretaceous calcareous nannofossils...Tanzania

the guidelines of Young et al. (1997). The biozonation isthat of Burnett (1998a). Only details of taxa and taxonomic references that do not appear in Bown (1998) are provided here. In the taxonomic descriptions below, ‘L’ =length, ‘W’ = width, ‘H’ = height. The taxonomic sectionis arranged in three parts: heterococcoliths, holococcoliths
and nannoliths. Within each part, taxa are arranged alphabetically, first according to the order, then family, thengenus, then species.
5.1 Heterococcoliths
Order ARKHANGELSKIALES Bown & Hampton in
Bown & Young, 1997
Family KAMPTNERIACEAE Bown & Hampton in
Bown & Young, 1997
Gartnerago chiasta Varol, 1991
Pl.3, figs 19, 20
Gartnerago? coxalliae sp. nov.
Pl.6, figs 34-37. Derivation of name: After Dr. Helen
Coxall (Cardiff University), Cenozoic micropalaeontologist on the TDP team. Diagnosis: A medium-sized species
with a rim that is reminiscent of that of Gartnerago(bright outer cycle, dark inner cycle, but lacking a central
plate). The wide central area is open and spanned by acomplex bar, aligned with the short axis, that remains
quite dark in XPL. The bar bifurcates where it joins therim. Differentiation: The bars in other species ofGartnerago are of simple construction and do not bifurcate. Holotype: Pl.6, figs 36, 37. Holotype dimensions:
L = 7.04µm, W = 4.84µm. Paratype: Pl.6, figs 34, 35.
Type locality: TDP5, Machole, south of Kitulo Hill, near
Lindi, coastal Tanzania. Type level: TDP5/10-2, 1516cm; UC15eTP, Late Campanian. Occurrence: TDP5
and TDP9, Late Campanian-Early Maastrichtian(UC15eTP-17); Matandu Bridge (Kilwa), Late Campanian(UC15eTP).
Gartnerago cf. G. praeobliquum Jakubowski, 1986
Pl.3, figs 21-24
Remarks: These specimens appear to differ from theholotype in having only simple arms making up the cross,rather than double, and in lacking clear ‘arrow-head’ terminations to the bars where they join the rim.
Order EIFFELLITHALES Rood et al., 1971
Family CHIASTOZYGACEAE Reinhardt, 1965
Amphizygus brooksii Bukry, 1969
Pl.6, figs 38-42
Chiastozygus spissus Bergen in Bralower & Bergen,
1998
Pl.9, figs 13-16

41

Gorkaea pseudanthophorus (Bramlette & Martini, 1964)
Varol & Girgis, 1994
Pl.7, figs 13-18
Placozygus banneri sp. nov.
Pl.7, figs 19-22. Derivation of name: After Prof. Fred
Banner (deceased), whose work on the planktonic
foraminifera of coastal Tanzania in the 1960s, with Prof.
Walter Blow, eventually led to the search for pristinepreservation that resulted in the TDP. Diagnosis: A small
species of Placozygus that has the spiral-rim extinction
pattern that is typical of the genus, a distinctive wide,
bright inner cycle, and a dark short-axis bar, apparently
composed of two blocks, that almost fills the small central
area. Differentiation: This new species is distinguishedfrom other placozygids by its wide, bright inner cycle,
narrow central area and almost indistinguishable shortaxis bar. Holotype: Pl.7, figs 21, 22. Holotype dimensions: L = 4.4µm, W = 3.08µm. Paratype: Pl.7, figs 19,
20. Type locality: Nguru-mahamba, SW of Lindi town,
coastal Tanzania. Type level: Sample Nguru-99-10;
UC15eTP, Late Campanian. Occurrence: Nguru-mahamba, Late Campanian-Early Maastrichtian (UC15eTP-17).
Placozygus fibuliformis (Reinhardt, 1964) Hoffmann,
1970
Pl.7, figs 27-30
Placozygus sp.
Pl.7, figs 31, 32
Placozygus spiralis (Bramlette & Martini, 1964)
Hoffmann, 1970
Pl.7, figs 23-26
Remarks: Burnett (1998a) has been somewhat responsible for confusing the placozygid taxonomy. The originaldescription of spiralis includes small size and small perforations either side of the bar; thus Burnett’s (1998a,
pl.6.2, fig.26a) illustration of P. fibuliformis is actually P.
spiralis. P. fibuliformis (illustrated here in Pl.7, figs 2730) has more open perforations and a bar in which the elements can be easily discerned. This species equates to
Burnett’s (1998a) illustrations (in pl.6.2, figs 26b-27b) of
P. cf. P. fibuliformis. The larger and more highly birefringent form illustrated by Burnett (1998a, pl.6.2, fig.28) as
P. spiralis, and here in Pl.7, figs 31 and 32 (as Placozygussp.), is possibly a new species.
Staurolithites ellipticus (Gartner, 1968) Lambert, 1987
Pl.8, figs 28-35
Staurolithites flavus Burnett, 1998b
Pl.8, figs 40-43
Staurolithites halfanii sp. nov.
Pl.6, figs 13-21. Derivation of name: After Mr. Halfani
R. Halfani (TPDC, Dar-es-Salaam), TDP facilitator.
Diagnosis: A small species of Staurolithites with an indistinct, unicyclic rim that possesses a dark, thin, axial cross

42

Lees

with a delicate spine and a small, distinctively highly birefringent spine-top, visible when the coccolith is rotatedaway from 0˚. Differentiation: This new species is distinguished by its bright spine-top and low-birefringencebars. Holotype: Pl.6, figs 19-21. Holotype dimensions: L= 3.96µm, W = 3.08µm. Paratypes: Pl.6, figs 13-18.
Type locality: TDP15, Maghreda Swamp, NW of Kilwa
Kivinje, coastal Tanzania. Type level: TDP15/31-1,
10cm; UC6b, Early Turonian. Occurrence: TDP15, Early
Turonian (UC6b).
Staurolithites handleyi sp. nov.
Pl.6, figs 28-33. Derivation of name: After Mr. Luke
Handley (Bristol University), organic geochemist on the
TDP team. Diagnosis: A small species of Staurolithites
with a bicyclic rim, the inner cycle of which is bright. The
axial cross appears to be composed of single bars that arebright at 0˚, becoming dark upon rotation.
Differentiation: This new species is distinguished by its
bicyclic rim and the particular simplicity of its cross.
Holotype: Pl.6, figs 30, 31. Holotype dimensions: L =3.96µm, W = 3.08µm. Paratypes: Pl.6, figs 28, 29, 32,
33. Type locality: TDP9, Nangurukuru junction, SW ofSingino Hill, near Kilwa Kivinje, coastal Tanzania. Typelevel: TDP9/14-1, 36cm; UC17, Early Maastrichtian.
Occurrence: TDP9, Late Campanian-Early Maastrichtian(UC15eTP-17).
Staurolithites ngurumahambaensis sp. nov.
Pl.6, figs 22-27. Derivation of name: From Ngurumahamba, where the species is described from.
Diagnosis: A small species of Staurolithites with a dark,
unicyclic rim, wherein the bars aligned with the long axisof the ellipse are dark, whilst the short-axis bars are highly birefringent. Differentiation: This new species is distinguished by its bright short-axis bars and dark long-axisbars. Holotype: Pl.6, figs 24, 25. Holotype dimensions:L = 3.08µm, W = 2.2µm. Paratypes: Pl.6, figs 22, 23, 26,
27. Type locality: Nguru-mahamba, SW of Lindi town,
coastal Tanzania. Type level: Sample Nguru-99-5; UC3ab, Middle-Late Cenomanian. Occurrence: Ngurumahamba, Middle-Late Cenomanian (UC3a-b); TDP15,
Early Turonian (UC6b).
Staurolithites zoensis Burnett, 1998b
Pl.8, figs 36-39
Zeugrhabdotus acanthus Reinhardt, 1965
Pl.7, figs 33-42; Pl.8, figs 1, 2
Zeugrhabdotus bicrescenticus (Stover, 1966) Burnett in
Gale et al., 1996
Pl.9, figs 1, 2
Zeugrhabdotus cf. Z. bicrescenticus (Stover, 1966)
Burnett in Gale et al., 1996
Pl.9, figs 3-7
Remarks: Z. cf. Z. bicrescenticus has two miniscule crystals, that are bright in XPL, inserted between the bars

where the bars join the rim, such as is seen in Z. diplogrammus. These are not apparent in Z. bicrescenticus.
Zeugrhabdotus biperforatus (Gartner, 1968) Burnett,
1998b
Pl.9, figs 8, 9
Zeugrhabdotus cf. Z. biperforatus (Gartner, 1968)
Burnett, 1998b
Pl.9, figs 10, 11
Remarks: Z. cf. Z. biperforatus is generally more narrowly elliptical, has a more-open central area and a narrower
inner cycle than Z. biperforatus.
Zeugrhabdotus blowii sp. nov.
Pl.8, figs 20-27. Derivation of name: After Prof. Walter
Blow (deceased), whose work on the planktonic
foraminifera of coastal Tanzania in the 1960s, with Prof.
Fred Banner, eventually led to the search for pristinepreservation that resulted in the TDP. Diagnosis: A small
species of Zeugrhabdotus with quite low birefringencethat has a simple, disjunct bar composed of two elements.
The bar is almost invisible at 0˚. At 45˚, the bar and rim
have the same birefringence. Differentiation: This new
species is distinguished from other zeugrhabdotids by thesimplicity of its bar and by the way in which the bar goes
into extinction at 0˚. The Jurassic Z. fissus is superficiallysimilar to the new species, but has a bar with higher birefringence and a perforation in the centre of the bar that the
new species lacks. Holotype: Pl.8, figs 22, 23. Holotype
dimensions: L = 3.52µm, W = 2.64µm. Paratypes: Pl.8,
figs 20, 21, 24, 25. Type locality: TDP5, Machole, southof Kitulo Hill, near Lindi, coastal Tanzania. Type level:
TDP5/10-2, 15-16cm; UC15eTP, Late Campanian.
Occurrence: TDP5, Late Campanian (UC15eTP); TDP15,
Early Turonian (UC6b).
Zeugrhabdotus embergeri (Noël, 1958) Perch-Nielsen,
1984
Pl.7, figs 1-12
Remarks: This heavily calcified species has a rhombshaped, composite bar and very small to no perforationsin the central area. The ragged-looking specimens in Pl.7,
figs 1-6 are interpreted as showing the proximal view ofthis species, with the rhomb-shaped base to the bar being
discernable, surrounded by the base-plate.
Zeugrhabdotus erectus (Deflandre in Deflandre & Fert,
1954) Reinhardt, 1965
Pl.9, figs 21-24; Pl.9, figs 19, 20?
Zeugrhabdotus noeliae Rood et al., 1971
Pl.8, figs 10-17
Zeugrhabdotus praesigmoides Burnett, 1998b
Pl.8, figs 3-9
Zeugrhabdotus simplex sp. nov.
Pl.8, figs 18, 19. Derivation of name: From the Latin

New Late Cretaceous calcareous nannofossils...Tanzania‘simplex’, meaning ‘plain’ or ‘simple’, referring to thesimplicity of the bar. Diagnosis: A large species ofZeugrhabdotus in which the slender, disjunct bar, which is
crystallographically uniform, divides the wide, open central area. Differentiation: This new species is distinguished from other zeugrhabdotids in having a simple,
slender bar dividing the wide, open central area.
Holotype: Pl.8, figs 18, 19. Holotype dimensions: L =8.80µm, W = 6.16µm. Type locality: TDP9,
Nangurukuru junction, SW of Singino Hill, near Kilwa
Kivinje, coastal Tanzania. Type level: TDP9/47-2, 30cm;
UC15eTP, Late Campanian. Occurrence: TDP9, Late
Campanian (UC15eTP).
Zeugrhabdotus xenotus (Stover, 1966) Burnett in Gale etal., 1996
Pl.9, figs 12, 17, 18
Family EIFFELLITHACEAE Reinhardt, 1965
Eiffellithus lindiensis sp. nov.
Pl.6, figs 1-12. Derivation of name: After the town ofLindi, near which the species was first identified.
Diagnosis: A small species of Eiffellithus in which thesmall, indistinct central cross is axially aligned. The innercycle fills the central area. Differentiation: This new
species is easily distinguished from E. eximius, which ismuch larger, and has a distinctive, bifurcating cross thattakes up more of the central area. Furthermore, E. lindiensis ranges up into the Maastrichtian. It is distinct fromother Late Cretaceous eiffellithids in being small and having an axial cross. Holotype: Pl.6, figs 3, 4. Holotype
dimensions: L = 4.4µm, W = 3.08µm. Paratypes: Pl.6,
figs 5-10. Type locality: TDP5, Machole, south of Kitulo
Hill, near Lindi, coastal Tanzania. Type level: TDP5/6-3,
36-37cm; UC15eTP, Late Campanian. Occurrence: TDP5
and TDP9, Late Campanian-Early Maastrichtian(UC15eTP-17); Lindi area, Campanian-Maastrichtian(UC16).
Family RHAGODISCACEAE Hay, 1977
Percivalia? dunkleyjonesii sp. nov.
Pl.3, figs 29-42. Derivation of name: After Mr. Tom
Dunkley Jones (UCL), Cenozoic nannopalaeontologist
and roustabout on the TDP team. Diagnosis: This medium-sized species possesses a narrow, bicyclic rim and acentral plate of laterally-oriented laths at the centre ofwhich is an indistinct cross. This axial cross has two distinctive ‘buttons’, outlined by dark extinction lines, making up the short-axis bars. The ‘buttons’ are visible when
the coccolith is oriented both at 0˚ and 45˚. The entire central area shows low birefringence. Differentiation: This
new species is distinguished from all other Percivalia
species by the distinctive ‘buttons’ and central-area laths.
Remarks: The rim and inner cycle of this species are
somewhat reminiscent of Percivalia, although scanning

43

electron microscopy is needed to confirm the genericassignment. Holotype: Pl.3, figs 40-42. Holotype dimensions: L = 6.82µm, W = 4.84µm. Paratypes: Pl.3, figs 3134, 37-39. Type locality: TDP15, Maghreda Swamp, NW
of Kilwa Kivinje, coastal Tanzania. Type level:
TDP15/25-1, 10cm; UC6b, Early Turonian. Occurrence:
TDP15, Early Turonian (UC6b); Nguru-mahamba, Early
Cenomanian (UC1a-2a).
Percivalia? pearsonii sp. nov.
Pl.3, figs 25-28. Derivation of name: After Prof. Paul
Pearson (Cardiff University), co-chief and Cenozoic
micropalaeontologist on the TDP team. Diagnosis: Thismedium-sized species possesses a narrow, bicyclic rim, acentral-area plate, and a central, axially-aligned cross,each bar of which is divided into two. The cross is located in the centre of a central plate. The cross and plateshow very low birefringence in XPL. Differentiation:
This new species is distinguished from other Percivalia
species by its distinctive axial cross and its lack of perforations in the central plate. Remarks: The rim, innercycle, and characteristically dark central area of this new
species are somewhat reminiscent of Percivalia, althoughscanning electron microscopy is needed to confirm thegeneric assignment. Holotype: Pl.3, figs 27, 28. Holotype
dimensions: L = 6.60µm, W = 4.40µm. Paratype: Pl.3,
figs 25, 26. Type locality: TDP9, Nangurukuru junction,
SW of Singino Hill, near Kilwa Kavinje, coastal
Tanzania. Type level: TDP9/21-1, 50cm; UC17, Early
Maastrichtian. Occurrence: TDP9 and Nguru-mahamba,
Early Maastrichtian (UC17).
Rhagodiscus pancostii sp. nov.
Pl.4, figs 1-4; Pl.4, figs 5, 6? Derivation of name: After
Dr. Richard Pancost (Bristol University), organic geochemist and palaeoceanographer on the TDP team.
Diagnosis: A small, elliptical species of Rhagodiscus thatbears a very small, delicate spine. The spine-tip appears
bright in XPL. Differentiation: Similar to R. achlyostaurion, but with a spine-base/top that is much smaller, anddoes not occupy most of the central area. R. gallagheri issomewhat similar, but has no spine and is more narrowly
elliptical. The specimen in Pl.4, figs 5, 6 has a narrower,
brighter rim, which differs from that seen in R. pancostii;
this specimen may be Orastrum partitum. Holotype: Pl.4,
figs 1, 2. Holotype dimensions: L = 3.96µm, W =2.86µm. Paratype: Pl.4, figs 3, 4. Type locality: TDP15,
Maghreda Swamp, NW of Kilwa Kivinje, coastal
Tanzania. Type level: TDP15/15-1, 10cm; UC6b, Early
Turonian. Occurrence: TDP15, Early Turonian (UC6b).
Order PODORHABDALES Rood et al., 1971, emend.
Bown, 1987
Family AXOPODORHABDACEAE Bown & Young,
1997

44

Lees

Cribrocorona echinus (Burnett, 1998b) Lees & Bown,
2005
Pl.4, figs 39-42
Cribrosphaerella circula (Risatti, 1973) comb. nov.
Pl.4, fig.37
Basionym: Cribrosphaera circula Risatti, 1973, p.24,
pl.8, figs 20, 21 (holotype). Risatti, J.B. 1973.
Nannoplankton biostratigraphy of the Upper Bluffport
Marl-Lower Prairie Bluff Chalk interval (Upper
Cretaceous), in Mississippi. In: L.A. Smith & J.
Hardenbol (Eds). Proceedings of Symposium on
Calcareous Nannofossils. SEPM, Gulf Coast section,
Houston, Texas, October 26 1973: 8-57.

whilst his light-micrographs (including the holotype)show specimens with a narrow central area.
Consequently, the form illustrated as M. ficula by Bergenin Bralower & Bergen (1998) is not ficula, but a new
species, bergenii. Holotype: Pl.5, figs 32-35. Holotype
dimensions: L = 6.82µm, W = 5.28µm. Paratypes: Pl.5,
figs 38-40. Type locality: Lindi area, coastal Tanzania.
Type level: Sample Lindi-99-11; UC17, Early
Maastrichtian. Occurrence: Lindi area, Early
Maastrichtian (UC17); TDP9, Late Campanian-Early
Maastrichtian (UC15eTP-17); TDP15, Early Turonian(UC6b); Bounds core, western Kansas (Bralower &Bergen, 1998), Middle-Late Turonian (UC8b-9b).

Cribrosphaerella ehrenbergii (Arkhangelsky, 1912)
Deflandre in Piveteau, 1952
Pl.4, figs 32-34 – typical subrectangular, coarse-grilledforms; Pl.4, figs 35, 36 – irregularly-rimmed forms; Pl.4,
fig.38 – elliptical, fine-grilled form

Family PREDISCOSPHAERACEAE Rood et al.,1971

Family BISCUTACEAE Black, 1971
Biscutum ellipticum (Górka, 1957) Grün in Grün &Allemann, 1975
Pl.5, figs 1-10, 12-15
Biscutum notaculum Wind & Wise in Wise & Wind,
1977
Pl.5, figs 18-21
Crucibiscutum hayi (Black, 1973) Jakubowski, 1986
Pl.5, figs 22, 23
Crucibiscutum salebrosum? (Black, 1971) Jakubowski,
1986
Pl.5, figs 24-31
Discorhabdus ignotus (Górka, 1957) Perch-Nielsen,
1968
Pl.5, figs 11, 16, 17
Family CRETARHABDACEAE Thierstein, 1973

Prediscosphaera mgayae sp. nov.
Pl.5, figs 36, 37, 41-44. Derivation of name: After Mr.
Elvis Mgaya (‘Mr. K’) (TPDC, Dar-es-Salaam), field
driver and drilling assistant on the TDP team. Diagnosis:
A small, elliptical species of Prediscosphaera with a delicate axial cross and a very narrow, low-birefringence
inner rim-cycle. Differentiation: P. spinosa has a thicker,
bright inner rim-cycle and a subrectangular outline. P.
stoveri is small, with a bright inner cycle that almost fills
the central area. P. arkhangelskyi has a distinctive crossand thick, bright inner cycle. Other species of the genus
have either circular outlines or crosses at 45˚ to the ellipse
axes. Holotype: Pl.5, figs 36, 37. Holotype dimensions:L = 4.84µm, W = 3.08µm. Paratypes: Pl.5, figs 41-44.
Type locality: TDP9, Nangurukuru junction, SW ofSingino Hill, near Kilwa Kivinje, coastal Tanzania. Typelevel: TDP9/19-1, 30cm; UC17, Early Maastrichtian.
Occurrence: TDP9 and Lindi area, Early Maastrichtian(UC17).
Order STEPHANOLITHIALES Bown & Young, 1997
Family CALCIOSOLENIACEAE Kamptner, 1927

Miravetesina bergenii sp. nov.
Non 1966 Coccolithites ficula Stover: p.138, pl.5, figs 5, 6; pl.9,
fig.11.
Non 1998b Retecapsa ficula (Stover) Burnett: pp.138-139; non
Burnett, 1998a, p.180, pl.6.7, fig.8.
1998 Miravetesina ficula (Stover, 1966) Bergen in Bralower &Bergen: p.75, pl.1, figs 13a, 13b; pl.2, fig.11.

Pl.5, figs 32-35, 38-40. Derivation of name: After Dr.
Jim Bergen (BP, Houston), seasoned calcareous nannofossil taxonomist and biostratigrapher. Diagnosis: A
medium-sized species of Miravetesina with a wide, granulate central area bearing a slightly sigmoidal, delicate,axial cross. Differentiation: Distinguished from M. favula (e.g. see Bown & Cooper, 1998, pl.4.13, fig.17) by its
more pronounced and slightly sigmoidal, axial cross, andits much younger age. Remarks: Retecapsa ficula, as
illustrated by Burnett (1998a), bears closest resemblance
to Stover’s (1966) line-drawing of ficula (pl.9, fig.11),

Calciosolenia fossilis (Deflandre in Deflandre & Fert,
1954) Bown in Kennedy et al., 2000
Pl.1, figs 1-6
Calciosolenia? huberi sp. nov.
Pl.1, figs 7-13; Pl.1, figs 14-18? Derivation of name:
After Dr. Brian Huber (Smithsonian, Washington),
Mesozoic micropalaeontologist on the TDP team.
Diagnosis: A medium-sized species of Calciosolenia?
possessing a distally-orientated spur-like extension to therim, located at one end of the coccolith, observable in sideview. Differentiation: No other species of Calciosolenia
possess a spur-like rim extension. This species is questionably placed in Calciosolenia based on a slightly
oblique view (Pl.1, fig.11) that suggests this assignment,
although it is unusual to observe Calciosolenia specimens
in side view. Specimens with thickened rims, shown on

New Late Cretaceous calcareous nannofossils...Tanzania

Pl.1, figs 14-18, may represent distal/proximal views ofthis species. Holotype: Pl.1, figs 9, 10. Holotype dimensions: L = 7.04µm, W = 1.1µm. Paratypes: Pl.1, figs 1113. Type locality: TDP15, Maghreda Swamp, NW of
Kilwa Kivinje, coastal Tanzania. Type level: TDP15/371, 10cm; UC6b, Early Turonian. Occurrence: TDP15,
Early Turonian (UC6b); TDP9, Early Maastrichtian(UC17).
Family STEPHANOLITHIACEAE Black, 1968
Corollithion completum Perch-Nielsen, 1973
Pl.1, figs 19, 20
Corollithion exiguum Stradner, 1961
Pl.1, figs 33, 34
Corollithion karegae sp. nov.
Pl.1, figs 25-32. Derivation of name: After Ms. Amina
Karega (TPDC, Dar-es-Salaam), nanno- andmicropalaeontologist on the TDP team. Diagnosis: A
small species of Corollithion with very low birefringence,a very narrow, slightly brighter inner rim-cycle, and analmost indistinguishable, dark outer cycle, in which aplate composed of six segments completely fills the central area. Differentiation: This new species has distinctively low birefringence. In other species, the bright innercycle is thicker and much more visible. The dark central
plate that fills the central area of the new species is composed of six segments, whilst in C. kennedyi (Pl.1, figs 21,22) and C. completum (Pl.1, figs 19, 20) the central plates
have higher birefringence and are divided into four
blocks. Holotype: Pl.1, figs 31, 32. Holotype dimensions: L = 3.96µm, W = 3.08µm. Paratype: Pl.1, figs 25,
26. Type locality: TDP5, Machole, south of Kitulo Hill,near Lindi, coastal Tanzania. Type level: TDP5/9-3, 5557cm; UC15eTP, Late Campanian. Occurrence: TDP5,
Late Campanian/Early Maastrichtian (UC15eTP-UC16);
TDP9, Late Campanian-Early Maastrichtian (UC15eTPUC17); Matandu Bridge (Kilwa), Late Campanian(UC15dTP); Lindi area, Campanian/Maastrichtian(UC16).
Corollithion kennedyi Crux, 1981
Pl.1, figs 21, 22
Corollithion signum Stradner, 1963
Pl.1, figs 23, 24
Darwinilithus pentarhethum Watkins in Watkins &Bowdler, 1984
Pl.4, figs 19-28
Rhombolithion rhombicum (Stradner & Adamiker, 1966)
Black, 1973
Pl.1, figs 35, 36
Rhombolithion rotatum (Rood et al., 1971) Black, 1973
Pl.1, figs 37, 38
Rotelapillus msakyae sp. nov.
Pl.4, figs 29-31. Derivation of name: After Ms. Emma45

Msaky (TPDC, Dar-es-Salaam), palynologist on the TDP
team. Diagnosis: A medium-sized species of Rotelapillus
that is elliptical, both in outline and central area. The central area contains a non-axially-aligned, low-angled crossthat is dark in XPL. Differentiation: This new species isdistinguished from all other cross-bearing Rotelapillus
species by its elliptical outline and central area; all otherspecies are circular in outline. Holotype: Pl.4, figs 29, 30.
Holotype dimensions: L = 6.16µm, W = 3.96µm.
Paratype: Pl.4, fig.31. Type locality: TDP9,
Nangurukuru junction, SW of Singino Hill, near Kilwa
Kivinje, coastal Tanzania. Type level: TDP9/9-2, 39cm;
UC17, Early Maastrichtian. Occurrence: TDP9, Early
Maastrichtian (UC17).
Truncatoscaphus macmillanii sp. nov.
Pl.1, figs 39-41. Derivation of name: After Dr. Ian
McMillan (Cardiff University), micropalaeontologist on
the TDP team. Diagnosis: A small, low-birefringencespecies of Truncatoscaphus with a relatively short andbroad polygonal rim, containing a clearly discernable longitudinal bar from which four very barely discernable,transverse bars extend down each side of the end thirds ofthe coccolith. Centrally, four further, clearly discernable
bars interlock to form a distinctive cross. Differentiation:
The new species most closely resembles the Jurassic T.
intermedius and T. delftensis, but both of these have moreelongated and narrower outlines, and the latter has morebars. Neither possesses the distinctively-arranged central
bars. The new species is the youngest Truncatoscaphus
recorded. Holotype: Pl.1, figs 39-41. Holotype dimensions: L = 3.52µm, W = 2.2µm. Type locality: TDP15,
Maghreda Swamp, NW of Kilwa Kivinje, coastal
Tanzania. Type level: TDP15/15-1, 10cm; UC6b, Early
Turonian. Occurrence: TDP15, Early Turonian (UC6b).
PLACOLITHS OF UNCERTAIN AFFINITY
Prolatipatella multicarinata Gartner, 1968
Pl.3, figs 13-16
Repagulum parvidentatum (Deflandre & Fert, 1954)
Forchheimer, 1972
Pl.3, figs 11, 12, 17, 18
5.2 Holococcoliths
Family CALYPTROSPHAERACEAE Boudreaux &Hay, 1969
Remarks: The classification of Mesozoic holococcoliths
is problematical. The majority are very small, therefore
rim-structure details and some central-area features are atthe very limits of light-microscopic observation. Becauseof their very low preservation potential, they are typicallyrare in occurrence, and so scanning electron microscopeobservation is often not practicable. Generic classificationis consequently subjective, being predominantly reliant
on central-area features. Additionally, matching side

46

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views of rare holococcoliths with their base-plate views is
not easy; most biostratigraphic analysis or reconnaissancework is done on permanently-mounted slides. This may
result in multiple taxonomic designations for the same
species.
Several previously unreported taxa are present in the
Tanzanian material. Whilst I attempted to match sideviews with base-plate views, the specimens are generallyrare and these images were taken on permanently-mounted smear-slides.
Acuturris scotus (Risatti, 1973) Wind & Wise in Wise &Wind, 1977
Pl.11, figs 22-24
Bifidalithus mchanae sp. nov.
Pl.10, figs 66-71. Derivation of name: After Mr. Ephrem
Mchana (TPDC, Dar-es-Salaam), for expert technicalassistance during TDP drilling. Diagnosis: A small
species of Bifidalithus Varol, 1991 in which the two main
constituent elements contain a single perforation each,arranged around the short-axis junction between the elements. The junction between these elements appears
raised. Differentiation: This species is distinct from B.
geminicatillus in possessing perforations. In side view
(Pl.10, figs 70, 71), the raised area where the elements
join appears to arch over a cavity above a monoperforate
proximal plate. Holotype: Pl.10, figs 68, 69. Holotype
dimensions: L = 4.62µm, W = 3.08µm. Paratypes: Pl.10,
figs 66, 67, 70, 71. Type locality: TDP5, Machole, southof Kitulo Hill, near Lindi, coastal Tanzania. Type level:
TDP5/9-1, 48-49cm; UC15eTP, Late Campanian.
Occurrence: TDP5, Late Campanian (UC15eTP); TDP9,
Early Maastrichtian (UC17).
Bilapillus gen. nov.
Type species: Bilapillus wadeae gen. et sp. nov.
Derivation of name: From the Latin ‘bi’, meaning ‘two’,and ‘lapillus’, meaning ‘little stone’, referring to the central structure of this genus. Diagnosis: A holococcolithwith a simple rim that is bright in XPL. At 0˚ in XPL,
extinction lines form v-shapes at the ends of the lith. On
rotation, the extinction lines become aligned with thepolarising directions, whilst the short-axis line is offset on
either side of the centre. Two adjacent polygonal blocks
fill the central area.
Bilapillus wadeae gen. et sp. nov.
Pl.13, figs 34-43; Pl.13, figs 44, 45? Derivation of name:
After Dr. Bridget Wade (Rutgers University), Cenozoic
micropalaeontologist on the TDP team. Diagnosis: A
small species of Bilapillus. The illustrations in Pl.13, figs
44, 45 appear to show perforations in the central blocks.
Holotype: Pl.13, figs 36, 37. Holotype dimensions: L =3.30µm, W = 2.20µm. Paratypes: Pl.13, figs 34, 35, 38,
39. Type locality: TDP15, Maghreda Swamp, NW of
Kilwa Kivinje, coastal Tanzania. Type level: TDP15/5-1,

10cm; UC6b, Early Turonian. Occurrence: TDP15, Early
Turonian (UC6b); Matandu Bridge (Kilwa), Middle-Late
Turonian (UC9a); Nguru-mahamba, Early Cenomanian(UC1a-2a).
Genus Calculites Prins & Sissingh in Sissingh, 1977
Remarks: New species that, in XPL, are bright across theentire holococcolith have here been placed in Calculites.
Calculites axosuturalis Bergen in Bralower & Bergen,
1998
Pl.9, figs 33-36, Pl.9, figs 37, 38?
Calculites cenomanicus sp. nov.
Pl.10, figs 37-42. Derivation of name: After the
Cenomanian, from which it is described. Diagnosis: A
small species of Calculites with an irregular-looking,elongated central perforation. At 0˚ in XPL, the extinctionlines are aligned with the polarising directions. On rotation, the extinction lines form v-shapes at the ends of thelith that are truncated by the ends of the perforation. The
central perforation appears to contain a structure (Pl.10,
figs 37, 38), but this is not resolvable. Differentiation:
The elongated, irregular-looking central perforation andvague central structure distinguish this from otherCalculites species. Holotype: Pl.10, figs 37, 38. Holotype
dimensions: L = 4.40µm, W = 2.64µm. Paratypes: Pl.10,
figs 39-42. Type locality: Nguru-mahamba, SW of Lindi
town, coastal Tanzania. Type level: Nguru-99-1; UC3a-b,
Middle-Late Cenomanian. Occurrence: Nguru-mahamba, Middle-Late Cenomanian (UC3a-b).
Calculites cyclops sp. nov.
Pl.10, figs 10-13; Pl.10, figs 14-17? Derivation of name:
From the Latin ‘Cyclops’, the mythical race of one-eyed
giants, referring to the single, relatively wide central perforation. Diagnosis: A small species of Calculites that hasa relatively wide central perforation. At 0˚ in XPL, theextinction lines are aligned with the polarising directions.
On rotation, the lines become v-shaped and occupy theends of the lith. Differentiation: The wide central perforation distinguishes it from other species of Calculites.
Holotype: Pl.10, figs 10, 11. Holotype dimensions: L =4.84µm, W = 3.52µm. Paratype: Pl.10, figs 12, 13. Type
locality: TDP15, Maghreda Swamp, NW of Kilwa
Kivinje, coastal Tanzania. Type level: TDP15/15-1,
10cm; UC6b, Early Turonian. Occurrence: TDP15, Early
Turonian (UC6b); ?Nguru-mahamba, Middle-Late
Cenomanian (UC3a-b).
Calculites juliae sp. nov.
Pl.13, figs 32, 33. Derivation of name: After Ms. Julia
Pearson, member of the TDP support team. Diagnosis: A
small species of Calculites, comprising a narrow rim
enclosing a multiperforate plate (six perforations, twolarger ones situated at each end of the lith and four smaller ones arranged around the outsides of the middle of theNew Late Cretaceous calcareous nannofossils...Tanzania

47

plate). The plate goes dark upon rotation. Differentiation:
The new species is smaller than most Calculites species
and has symmetrically-distributed perforations around thecentral plate, which distinguishes it from other species.
Holotype: Pl.13, figs 32, 33. Holotype dimensions: L =3.30µm, 2.20µm. Type locality: Nguru-mahamba, SW of
Lindi town, coastal Tanzania. Type level: Nguru-99-9;
UC17, Early Maastrichtian. Occurrence: Nguru-mahamba, Early Maastrichtian (UC17).

the extinction lines are aligned with the polarising directions. On rotation, the extinction lines join with the central slit to form an H-shape. Differentiation: The central
slit distinguishes this from other Calculites species.
Holotype: Pl.10, figs 43, 44. Holotype dimensions: L =4.18µm, W = 2.86µm. Type locality: TDP15, Maghreda
Swamp, NW of Kilwa Kivinje, coastal Tanzania. Typelevel: TDP15/25-1, 10cm; UC6b, Early Turonian.
Occurrence: TDP15, Early Turonian (UC6b).

Calculites maghredaswampensis sp. nov.
Pl.10, figs 19-22; Pl.10, figs 23-26? Derivation of name:
After Maghreda Swamp, where TDP15 was drilled.
Diagnosis: A medium-sized species of Calculites with asmall central perforation. In XPL at 0˚, the extinctionlines are are aligned with the polarising directions, but
kink as they move into the central area, where they meetin the middle. On rotation, these form a sigmoidal x-shapeacross the lith. Differentiation: Distinguished from otherCalculites by its kinked extinction lines and central perforation. Holotype: Pl.10, figs 19, 20. Holotype dimensions: L = 5.28µm, W = 3.52µm. Paratype: Pl.10, figs
21, 22. Type locality: TDP15, Maghreda Swamp, NW of
Kilwa Kivinje, coastal Tanzania. Type level: TDP15/7-1,
10cm; UC6b, Early Turonian. Occurrence: TDP15, Early
Turonian (UC6b); ?TDP5, Late Campanian (UC15eTP).

Calculites rosalyniae sp. nov.
Pl.13, figs 30, 31. Derivation of name: After Ms.
Rosalyn Pearson, member of the TDP support team.
Diagnosis: A small species of Calculites comprising four
blocks divided by wavy sutures that form an H-shape.
Each plate has a perforation towards the centre. The
blocks appear to go into extinction separately. The lithappears similar at all angles of rotation in XPL.
Differentiation: The new species is superficially similar
to C. obscurus, but bears a perforation in each quadrant,
and the extinction pattern is dissimilar. Holotype: Pl.13,
figs 30, 31. Holotype dimensions: L = 4.40µm, W =3.08µm. Type locality: TDP5, Machole, south of Kitulo
Hill, near Lindi, coastal Tanzania. Type level: TDP5/5-1,
LateCampanian-Early
66-67cm;
UC15eTP-16,
Maastrichtian. Occurrence: TDP5, Late CampanianEarly Maastrichtian (UC15eTP-16).

Calculites obscurus (Deflandre, 1959) Prins & Sissinghin Sissingh, 1977
Pl.9, figs 29-32
Calculites paulus sp. nov.
Pl.10, figs 18, 27. Derivation of name: From the Latin‘paulus’, meaning ‘little’, referring to the size of thisspecies. Diagnosis: A very small species of Calculites thathas an elongate, narrow central perforation. At 0˚ in XPL,
two extinction lines are aligned with the polarising directions, but the short-axis line is offset. On rotation, thelines become v-shaped and occupy the ends of the lith.
Differentiation: The very small size and elongate, narrow
central perforation distinguish this species from otherspecies of Calculites. Holotype: Pl.10, figs 18, 27.
Holotype dimensions: L = 2.64µm, W = 1.76µm. Type
locality: TDP9, Nangurukuru junction, SW of Singino
Hill, near Kilwa Kivinje, coastal Tanzania. Type level:
TDP9/9-2, 39cm; UC17, Early Maastrichtian.
Occurrence: TDP9, Early Maastrichtian (UC17).
Calculites percernis Jeremiah, 1996
Pl.10, figs 46-53
Calculites proscissus sp. nov.
Pl.10, figs 43, 44; Pl.10, figs 45, 54? Derivation of name:
From the Latin ‘proscissus’, meaning ‘slit’, referring to
the central slit-like perforation typical of this species.
Diagnosis: A small species of Calculites with an elongated, very narrow central perforation (or slit). At 0˚ in XPL,

Calculites turonicus sp. nov.
Pl.10, figs 33-36. Derivation of name: After theTuronian, from which it is described. Diagnosis: A small
species of Calculites which bears a very small central perforation and in which the central part is slightly darker in
XPL than the outer part (there is no discernable rim). At
0˚ in XPL, the extinction lines are aligned with the polarising directions and bifurcate (indicating a rim block in
extinction?) at each end of the lith. The central perforation
is not obvious in this direction, where the extinction lines
meet. On rotation, the extinction lines form v-shapes that
do not join in the centre of the lith. The small perforationis then clear. Differentiation: The new species bears aresemblance to C. percernis (see Pl.10, figs 46-53), but
the latter is generally smaller and the central area is notslightly darker than the rim. The extinction lines of C.
percernis are also different, with the short axis line being
kinked, so as to appear offset, rather than straight, as in
the new species. Holotype: Pl.10, figs 33, 34. Holotype
dimensions: L = 3.96µm, W = 2.86µm. Paratype: Pl.10,
figs 35, 36. Type locality: TDP15, Maghreda Swamp,
NW of Kilwa Kivinje, coastal Tanzania. Type level:
TDP15/25-1, 10cm; UC6b, Early Turonian. Occurrence:
TDP15, Early Turonian (UC6b).
Duocameratus gen. nov.
Type species: Duocameratus leariae gen. et sp. nov.
Derivation of name: From the Latin ‘duo’, meaning two,‘camera’, meaning ‘vaulted chamber’ and ‘atus’, mean-

48

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ing ‘like’, referring to the double-chambered, vaulted
nature of the holococcolith in side view. Diagnosis: In
side view, delicate-looking, cavate, polygonal holococcoliths with thin rims and proximal plates. A central septum
divides the cavity into two. Where the septum meets theexternal structure, the joints are thickened and appear
bright in most orientations.
Duocameratus leariae gen et sp. nov.
Pl.13, figs 51-59. Derivation of name: After Dr. Carrie
Lear, Cenozoic palaeoceanographer on the TDP team.
Diagnosis: A small species of Duocameratus in which the
distal(?) end of the septum is brightest and thickest, forming a subtriangular apex that distinguishes this species.
Holotype: Pl.13, figs 51-53. Holotype dimensions: H =4.40µm, W = 3.52µm. Paratype: Pl.13, figs 54-57. Type
locality: Nguru-mahamba, SW of Lindi town, coastal
Tanzania. Type level: Nguru-99-1; UC3a-b, Middle-Late
Cenomanian. Occurrence: Nguru-mahamba, MiddleLate Cenomanian (UC3a-b).
Duocameratus sianiae gen et sp. nov.
Pl.13, figs 60-65. Derivation of name: After Dr. Siani
Pearson (Cardiff), member of the TDP support team.
Diagnosis: A small species of Duocameratus that is irregularly pentagonal and house-shaped, with an almost flatbase/proximal plate and a pointed ‘roof’. Bright points
mark where the septum dividing the cavity joins the rim.
These bright points are situated in the middle of the baseand the peak of the ‘roof’. Holotype: Pl.13, figs 60, 61.
Holotype dimensions: H = 3.08µm, W = 3.52µm.
Paratypes: Pl.13, figs 62-65. Type locality: TDP15,
Maghreda Swamp, NW of Kilwa Kivinje, coastal
Tanzania. Type level: TDP15/5-1, 10cm; UC6b, Early
Turonian. Occurrence: TDP15, Early Turonian (UC6b).
Holococcolith indeterminate 1 side view
Pl.11, figs 47-49
Holococcolith indeterminate 2 side view
Pl.11, figs 50-52
Holococcolith indeterminate 3 side view
Pl.13, figs 48-50
Holococcolith? indeterminate side view
Pl.13, figs 66-68
Genus Lucianorhabdus Deflandre, 1959
Remarks: New species that have significant spines thatare most frequently observed in side view are here placedin Lucianorhabdus.
Lucianorhabdus? boudagherfadeliae sp. nov.
Pl.11, figs 43-46. Derivation of name: After Dr. Marcelle
Boudagher-Fadel (UCL), planktonic and larger benthic
foraminifera expert. Diagnosis: A small species ofLucianorhabdus? with a relatively short, hollow spinethat terminates bluntly and at an angle. The angled termination is visible when the lith is rotated, in XPL. The

small base-plate rim is relatively thick and the plate has athin proximal cycle. The base-plate is perforate and contains a structure that is not discernable. Differentiation:
Other lucianorhabdids do not possess the central structurethat is hinted at as lying beneath the spine-base. The shortspine and its angled termination is distinctive. Holotype:
Pl.11, figs 43-46. Holotype dimensions: H = 4.40µm, W= 4.40µm. Paratype: Pl.11, figs 43, 44. Type locality:
Nguru-mahamba, SW of Lindi town, coastal Tanzania.
Type level: Nguru-99-1; UC3a-b, Middle-Late
Cenomanian. Occurrence: Nguru-mahamba, MiddleLate Cenomanian (UC3a-b).
Lucianorhabdus compactus (Verbeek, 1976) comb. nov.
Basionym: Isocrystallithus compactus Verbeek, 1976,
p.78, pl.2, figs 1-4 (3 = holotype). Verbeek, J.W. 1976.
Upper Cretaceous calcareous nannoplankton from Ballon
and Théligny in the type area of the Cenomanian stage(Sarthe, France). Proceedings of the Koninklijke
Nederlandse Akademie van Wetenschappen, B79: 69-82.
Lucianorhabdus maleformis Reinhardt, 1966
Pl.11, figs 17, 18
Lucianorhabdus? tabernus sp. nov.
Pl.12, figs 45-47. Derivation of name: From the Latin‘taberna’, meaning ‘hut’, referring to the simple hutshaped outline of the species. Diagnosis: A cavate speciesof Lucianorhabdus? with a medium-sized base-plate andshort, conical spine. In XPL at 0˚, the lith is of low birefringence, and the thin base-plate is almost invisible. On
rotation, the spine becomes bright and the base-plate justvisible. Differentiation: Shorter than other lucianorhabdids, with a wider spine and equilateral-triangular outline.
The new species bears some resemblance to the
Paleocene Semihololithus tentorium Bown, 2005 (pl.31,
figs 11-20), however, the new species has a much thinner,
simpler base-plate and goes completely into extinction at
0˚, unlike S. tentorium. Holotype: Pl.12, figs 45-47.
Holotype dimensions: H = 5.28µm, W = 5.72µm. Type
locality: Nguru-mahamba, SW of Lindi town, coastal
Tanzania. Type level: Nguru-99-5; UC3a-b, Middle-Late
Cenomanian. Occurrence: Nguru-mahamba, MiddleLate Cenomanian (UC3a-b).
Lucianorhabdus turris sp. nov.
Pl.11, figs 1-6. Derivation of name: From the Latin ‘turris’, meaning ‘tower’, referring to the solid base and long,tapering spine of the species. Diagnosis: A species ofLucianorhabdus with a relatively long, hollow spine, thattapers to a point. The spine may be straight or may lean toone side. In XPL, the medium-sized base-plate has a narrow rim and appears as a solid block in the middle whenrotated, whilst at 0˚, there appears to be a central perforation in the plate, as it becomes optically discontinuous.
Differentiation: The new species differs from otherlucianorhabdids in having a pointed spine, that may orNew Late Cretaceous calcareous nannofossils...Tanzania

may not lean, without a terminal plug or plate. It is distinguished from Acuturris scotus by its larger size and thick,solid base-plate. Holotype: Pl.11, figs 1-4. Holotype
dimensions: H = 8.36µm, W = 5.72µm. Paratype: Pl.11,
figs 5, 6. Type locality: Nguru-mahamba, SW of Lindi
town, coastal Tanzania. Type level: Nguru-99-5; UC3a-b,
Middle-Late Cenomanian. Occurrence: Nguru-mahamba, Middle-Late Cenomanian (UC3a-b).
Metadoga ampulla Lees & Varol sp. nov.
Pl.11, figs 25-28. Derivation of name: From the Latin‘ampulla’, meaning ‘flask’, referring to the similarity of
the coccolith in side view to a standard laboratory flask.
Diagnosis: A small species of Metadoga that has a simple
flask-shaped outline in side-view. The holococcolith iscavate and the upper half of the lith goes into extinctionin XPL at 0˚. Differentiation: The new species differsfrom other species of Metadoga in having the distinctive
standard flask outline and in lacking a mid-lith collar.
Holotype: Pl.11, figs 25, 26. Holotype dimensions: H =4.40µm, W = 4.18µm. Type locality: TDP5, Machole,south of Kitulo Hill, near Lindi, coastal Tanzania. Typelevel: TDP5/9-1, 48-49cm; UC15eTP, Late Campanian.
Occurrence: TDP5, Late Campanian (UC15eTP); Ngurumahamba, Middle-Late Cenomanian (UC3a-b); Arabian
Peninsula, Early Campanian (Varol, pers. obs., 2007).
Munarinus keadyi Risatti, 1973
Pl.13, figs 14, 15, 18-25; Pl.13, figs 16, 17 side view?
Munarinus lesliae Risatti, 1973
Pl.13, figs 10-13
Munarinus mkeremei sp. nov.
Pl.13, figs 8, 9. Derivation of name: After Mr. Michael
Mkereme (TPDC, Dar-es-Salaam), for expert technicalassistance during TDP drilling. Diagnosis: A small
species of Munarinus possessing a relatively wide central
perforation that is spanned by a disjunct bar that joins therim with straight edges and spans the short axis. At 0˚ in
XPL, the extinction lines are aligned with the polarising
directions, and on rotation these form v-shapes at the endsof the lith. The bar is not visible at 0˚, becoming bright onrotation. Differentiation: M. keadyi is smaller, with a narrower central perforation and with a short-axis bar that is
subrhombic. The pointed ends of that bar slot into the rim.
Holotype: Pl.13, figs 8, 9. Holotype dimensions: L =4.84µm, W = 3.52µm. Type locality: TDP5, Machole,south of Kitulo Hill, near Lindi, coastal Tanzania. Typelevel: TDP5/9-1, 48-49cm; UC15eTP, Late Campanian.
Occurrence: TDP5, Late Campanian (UC15eTP).
Nicholasia gen. nov.
Type species: Nicholasia baileyi gen. et sp. nov.
Derivation of name: After Dr. Chris Nicholas (Trinity
College, Dublin University), sedimentologist, structuralgeologist and TDP co-chief. Diagnosis: Delicate-looking
holococcoliths that, in side view, have the outline of a hol-49

low isosceles triangle. The base-plate is virtually invisible. Where the spine joins the base-plate, there are twobright spots.
Nicholasia baileyi gen et sp. nov.
Pl.13, figs 69-73. Derivation of name: After Dr. Trevor
Bailey (Natural History Museum of Wales, Cardiff),
Cenozoic palaeoceanographer on the TDP team.
Diagnosis: As for the genus. This species is very small.
The ‘spine’ appears to terminate bluntly and is generally
dark. Holotype: Pl.13, fig.71. Holotype dimensions: H =2.20µm, W = 1.76µm. Paratypes: Pl.13, figs 69, 70, 72,
73. Type locality: Nguru-mahamba, SW of Lindi town,
coastal Tanzania. Type level: Nguru-99-10; UC15eTP,
Late Campanian. Occurrence: Nguru-mahamba, Late
Campanian (UC15eTP); TDP15, Early Turonian (UC6b);
Matandu Bridge (Kilwa), Middle-Late Turonian (UC9a).
Genus Orastrum Wind & Wise in Wise & Wind, 1977
Remarks: New species with bright rims and dark central
plates have here been placed in Orastrum.
Orastrum perspicuum Varol in Al-Rifaiy et al., 1990
Pl.10, figs 55-58 – nonperforate forms; Pl.10, figs 59, 60- monoperforate form; Pl.10, figs 61, 62 – monoperforate form?; Pl.10, figs 63-65, 72 – biperforate forms
Orastrum robinsonii sp. nov.
Pl.10, figs 1-9. Derivation of name: After Dr. Stuart
Robinson (UCL), Cretaceous palaeoenvironmental specialist. Diagnosis: A medium-sized species of Orastrum
that in XPL has a bright rim and dark central plate, bearing a usually barely-discernable x-shaped structure. At 0˚
in XPL, the extinction lines are aligned with the polarising directions, appearing darker on the rim and barely discernable on the plate. On rotation, these become v-shapedon the rim, occupying the ends of the lith, but bend where
they move onto the plate, where again they are barely discernable. In PC, the plate appears to be multiperforate(Pl.10, fig.3). Differentiation: The size and relatively
wide rim and central area are distinctive. The x-shaped
central structure distinguishes it from other species of thegenus. Holotype: Pl.10, figs 4, 5. Holotype dimensions:
L = 6.16µm, 4.84µm. Paratypes: Pl.10, figs 1-3, 6, 7.
Type locality: Nguru-mahamba, SW of Lindi town,
coastal Tanzania. Type level: Nguru-99-1; UC3a-b,
Middle-Late Cenomanian. Occurrence: Nguru-mahamba, Early-Late Cenomanian (UC1a-3b).
Ottavianus terrazetus Risatti, 1973
Pl.13, figs 1-7
Owenia dispar? (Varol in Al-Rifaiy et al., 1990) Bown
in Kennedy et al., 2000
Pl.12, figs 14-17
Owenia hillii Crux, 1991
Pl.12, figs 1-8, 10-13; Pl.12, figs 19-36? side views

Lees

50

Owenia cf. O. hillii? Crux, 1991 side view
Pl.12, figs 48, 49
Remarks: Unlike O. hillii side view, O. cf. O. hillii? sideview has a much shorter spine that either flares at the end,
or is capped by a plug that overhangs the top of the spine.
Pearsonia gen. nov.
Type species: Pearsonia ecclesiata gen. et sp. nov.
Derivation of name: After Prof. Paul Pearson (Cardiff
University), TDP co-chief and Cenozoic micropalaeontologist on the TDP team. Diagnosis: Cavate holococcoliths
with thin rims and small spines, which act as a single
crystal in XPL.
Pearsonia ecclesiata gen et sp. nov.
Pl.11, figs 29-42. Derivation of name: From the Latin‘ecclesia’, meaning ‘church’, and ‘atus’, meaning ‘likeness to’, referring to its resemblance to a church with aspire in side view. Diagnosis: A very small species ofPearsonia with a church-plus-spire-shaped outline in sideview. At 0˚, the liths are dark in XPL, becoming brightwhen rotated, with the base-plate and spine behaving as asingle crystal. Holotype: Pl.11, figs 29-32. Holotype
dimensions: H = 3.08µm, L = 2.64µm. Paratype: Pl.11,
figs 34, 35. Type locality: Matandu Bridge, west of Kilwa
Kivinje, coastal Tanzania. Type level: PP04-K1; UC9a,
Middle-Late Turonian. Occurrence: Matandu Bridge(Kilwa), Middle-Late Turonian (UC9a); TDP15, Early
Turonian (UC6b).
Petrobrasiella venata Troelsen & Quadros, 1971
Pl.9, figs 25-28
Russellia bukryi Risatti, 1973
Pl.10, figs 28-32
Russellia laswellii Risatti, 1973
Pl.13, figs 26-29
Tanzanella gen. nov.
Type species: Tanzanella bownii gen. et sp. nov.
Derivation of name: After Tanzania, where the genus isdescribed from. Diagnosis: Holococcoliths with a complex structure, as seen in side view. In XPL, rotated to 45˚,
the rim and proximal plate are thin, birefringent and clearly visible. Encased within these is a blocky structure, fromthe distal surface of which emanate four ‘prongs’.
Tanzanella bownii gen et sp. nov.
Pl.12, figs 37-44. Derivation of name: After Dr. Paul
Bown (UCL) in recognition of his sterling Cenozoic calcareous nannofossil taxonomic work on the TDP material. Diagnosis: A medium-sized species of Tanzanella.
Holotype: Pl.12, figs 43, 44. Holotype dimensions: L =5.72µm, H = 2.64µm. Paratypes: Pl.12, figs 37-42. Type
locality: Nguru-mahamba, SW of Lindi town, coastal
Tanzania. Type level: Nguru-99-1; UC3a-b, Middle-Late
Cenomanian. Occurrence: Nguru-mahamba, MiddleLate Cenomanian (UC3a-b).

Thecatus gen. nov.
Type species: Thecatus varolii gen. et sp. nov. Derivationof name: From the Latin ‘theca’, meaning ‘box’, and‘atus’ meaning ‘like’, referring to the boxy, cavate shapeof this holococcolith in size view. Diagnosis: A cavate,box-shaped holococcolith. At 0˚, in side view in XPL, therim is relatively tall and the proximal plate thin. A bipartite structure/short spine sits atop the rim, completing thebox shape. The spine is not visible on rotation.
Thecatus varolii gen. et sp. nov.
Pl.13, figs 46, 47. Derivation of name: After Dr. Osman
Varol (Varol Research, Llandudno), seasoned calcareous
nannofossil biostratigrapher and taxonomist. Diagnosis:
As for the genus. This species is small. Holotype: Pl.13,
figs 46, 47. Holotype dimensions: H = 3.08µm, W =3.52µm. Type locality: TDP15, Maghreda Swamp, NW
of Kilwa Kivinje, coastal Tanzania. Type level:
TDP15/29-1, 10cm; UC6b, Early Turonian. Occurrence:
TDP15, Early Turonian (UC6b).
5.3 Nannoliths
Family BRAARUDOSPHAERACEAE Deflandre,
1947
Braarudosphaera bigelowii (Gran & Braarud, 1935)
Deflandre, 1947
Pl.2, fig.28
Goniolithus fluckigeri Deflandre, 1957
Pl.2, figs 29, 30
Micrantholithus? nicholasii sp. nov.
Pl.2, figs 23-27. Derivation of name: After Dr. Chris
Nicholas (Trinity, Dublin University), co-chief, generalgeologist and lithostratigrapher on the TDP team.
Diagnosis: Possible segments of pentaliths ofMicrantholithus, each segment having a median depression, bordered by ridges, occupying the central third of
the segment. The expression of this depression at theouter edge of the segment creates a very shallow ‘w’shape. Differentiation: This new species is composed of
distinctively-shaped segments with a ridge-bordered central depression and crenulated outer edge that has not been
seen in any other species. Remarks: Although no entire
pentaliths have yet been found, the segment angles suggest a pentalith structure, and so this new species is questionably placed into Micrantholithus. Holotype: Pl.2,
fig.27. Holotype dimensions: Central lith point to outsideedge = 4.40µm, point to point across outside edge =3.96µm. Paratype: Pl.2, fig.25. Type locality: TDP15,
Maghreda Swamp, NW of Kilwa Kivinje, coastal
Tanzania. Type level: TDP15/15-1, 10cm; UC6b, Early
Turonian. Occurrence: TDP15, Early Turonian (UC6b).

New Late Cretaceous calcareous nannofossils...Tanzania

Family MICRORHABDULACEAE Deflandre, 1963
Lithraphidites acutus Verbeek & Manivit in Manivit etal., 1977
Pl.2, figs 21, 22
Lithraphidites carniolensis Deflandre, 1963
Pl.2, figs 1-3; Pl.2, figs 4-7 – thick form
Rhabdolithus aquitanicus Manivit, 1971
Pl.2, figs 8-20
Family INCERTAE SEDIS‘Anacanthoica mitra’ Varol, 1989
Pl.12, figs 9, 18
Remarks: As Bown (2005, p.38) remarked, these forms
may be fragments of a calcisphere test.
Ceratolithina capitanea (Burnett, 1997a) stat. nov.
Pl.2, figs 31-34
1997a Ceratolithina cruxii Perch-Nielsen, 1988 subsp. capitanea Burnett: p.59, pl.1, fig.4 (holotype).

Basionym: Ceratolithina cruxii Perch-Nielsen, 1988
subsp. capitanea Burnett, 1997a. Burnett, J.A. 1997a.
‘Middle’ Cretaceous morphological diversity within thegenus Ceratolithina Martini, 1967. Journal of
Nannoplankton Research, 19(1): 57-65. Remarks: This
species is clearly distinct from C. cruxii, having a much
longer horn that is concave along the edges, rather thanstraight or convex, as in C. cruxii. Consequently, capitanea is herein elevated to species status.
Ceratolithoides ohalloraniae sp. nov.
Pl.2, figs 35-40. Derivation of name: After Ms. Aoife
O’Halloran (Trinity, Dublin University), sedimentologist/clay mineralogist and geochemist on the TDP team.
Diagnosis: A small species of Ceratolithoides with analmost equilateral-triangular outline and a shallow interhorn angle. The cone extends halfway down the length of
the nannolith and is bright at 45˚. Differentiation: This
new species is smaller than the majority of
Ceratolithoides species. Its almost equilateral-triangularoutline distinguishes it from all other species. Holotype:
Pl.2, figs 39, 40. Holotype dimensions: Cone-tip to interhorn angle = 2.2µm, side and horn-point to horn-point =3.08µm. Paratype: Pl.2, figs 35, 36. Type locality:
TDP9, Nangurukuru junction, SW of Singino Hill, near
Kilwa Kivinje, coastal Tanzania. Type level: TDP9/9-2,
39cm; UC17, Early Maastrichtian. Occurrence: TDP9and Lindi area, Early Maastrichtian (UC17).
Ceratolithoides prominens? Burnett, 1997b
Pl.3, figs 1-4
Remarks: C. prominens is distinguished by its prominentcone, isosceles-triangular shape, short internal cone
length, and obtuse inter-horn angle. The specimens illustrated here are similar but with less prominent cones, possibly because they are etched specimens.

51

Ceratolithoides dongenii sp. nov.
Pl.3, figs 5-10. Derivation of name: After Dr. Bart van
Dongen (Stockholm University), organic geochemist on
the TDP team. Diagnosis: A medium-sized species of
Ceratolithoides with an almost equilateral-triangular outline, a prominent cone, almost entirely enclosed by thehorns, occupying half the length of the ceratolith, and aninter-horn angle approaching 90˚. Differentiation: This
new species has a shorter, broader cone (e.g. Pl.3, fig.5)
and a more equilateral-triangular shape than C. aculeus. It
somewhat resembles C. prominens, but the cone is entirely enclosed by the horns, the outline is triangular, rather
than subtriangular, the inter-horn angle is more acute, andthe cone is less prominent in the new species. The cone in
C. sagittatus Lees & Bown, 2005 is not enclosed by thehorns, and the cone divides into two elements, which distinguishes this from the new species. Holotype: Pl.3, figs
7, 8. Holotype dimensions: Cone-tip to inter-horn angle= 3.08µm, side = 5.06µm, horn-point to horn-point =4.40µm. Paratype: Pl.3, figs 9, 10. Type locality: TDP5,
Machole, south of Kitulo Hill, near Lindi, coastal
Tanzania. Type level: TDP5/9-3, 55-57cm; UC15eTP,
Late Campanian. Occurrence: TDP5, Late Campanian(UC15eTP); TDP9, Early Maastrichtian (UC17); Kitulo
Hill (Lindi), Early Maastrichtian (UC18).
Singanoa gen. nov.
Type species: Singanoa scapus gen. et sp. nov.
Derivation of name: After Dr. Joyce Singano (TPDC,
Dar-es-Salaam), micropalaeontologist, general facilitatorand ‘mother’ of the TDP team. Diagnosis: Nannolith ofvarious lengths, comprising four narrow blocks, arranged
end-to-end in two pairs. Diagonal blocks go into extinction together, and a dark extinction area in the centre gives
the appearance of a constriction in the nannolith (like ascroll tied at the middle).
Singanoa scapus gen. et sp. nov.
Pl.4, figs 7-18. Derivation of name: From the Latin ‘scapus’, meaning ‘a scroll’, referring to the scroll-likeappearance of the nannolith. Diagnosis: As for the genus.
May be medium-sized to very small. Differentiation: The
short form of this species bears a slight resemblance to thebirefringent blades of Lithraphidites? charactozorro SelfTrail, 1999, as seen in XPL. S. scapus is distinct, however, in not having the tapering ends possessed by thatspecies, and in being much smaller. Holotype: Pl.4, figs
11, 12. Holotype dimensions: L = 6.16µm, W = 0.88µm.
Paratype: Pl.4, figs 7-10, 13-18. Type locality: TDP9,
Nangurukuru junction, SW of Singino Hill, near Kilwa
Kivinje, coastal Tanzania. Type level: TDP9/15-1, 21cm;
UC17, Early Maastrichtian. Occurrence: TDP9, Late
Campanian-Early Maastrichtian (UC15eTP-UC17).
Acknowledgements
The author is very grateful to the NERC for funding (Grant
NE/C510508/1), and also to the TDP team, especially Dr. Joyce

52

Lees

Singano (TPDC), Prof. Paul Pearson (Cardiff University) and
Dr. Chris Nicholas (Trinity College, Dublin University), forbeing a joy to work with. Prof. Paul Pearson is especiallythanked for inviting me to be involved. As ever, Dr. Paul Bown
(UCL) was indispensable, especially as the manuscript neared
completion. Finally, many, many thanks to the reviewers: Drs
Osman Varol (Varol Research), Jean Self-Trail (USGS) and Paul
Bown; you all accomplished a heroic feat in reviewing thismuch taxonomy so quickly for me, and I certainly owe you.

References
Bown, P.R. (Ed.) 1998. Calcareous Nannofossil
Biostratigraphy. British Micropalaeontological Society
Publications Series. Chapman & Hall/Kluwer Academic
Publishers, London: 315pp.
Bown, P.R. 2005. Palaeogene calcareous nannofossils from theKilwa and Lindi areas of coastal Tanzania (Tanzania
Drilling Project 2003-4). Journal of Nannoplankton
Research, 27: 21-95.
Bown, P.R. & Cooper, M.K.E. 1998. Jurassic. In: P.R. Bown
(Ed.). Calcareous Nannofossil Biostratigraphy. British
Micropalaeontological Society Publications Series.
Chapman & Hall/Kluwer Academic Publishers, London: 3485.
Bown, P.R. & Dunkley Jones, T. 2006. New Palaeogene calcareous nannofossil taxa from coastal Tanzania: Tanzania
Drilling Project Sites 11 to 14. Journal of Nannoplankton
Research, 28: 17-34.
Bown, P.R., Dunkley Jones, T. & Young, J.R. 2007.
Umbilicosphaera jordanii Bown, 2005 from the Paleogeneof Tanzania: confirmation of generic assignment and a
Paleocene origination for the Family Calcidiscaceae.
Journal of Nannoplankton Research, 29: 25-30.
Bown, P.R., Lees, J.A. & Young, J.R. 2004. Calcareous nannoplankton evolution and diversity through time. In: H.R.
Thierstein & J.R. Young (Eds). Coccolithophores - From
Molecular Processes to Global Impact. Springer, Berlin:481-508.
Bown, P.R. & Young, J.R. 1998. Techniques. In: P.R. Bown
(Ed.). Calcareous Nannofossil Biostratigraphy. British
Micropalaeontological Society Publications Series.
Chapman & Hall/Kluwer Academic Publishers, London: 1628.
Bralower, T.J. & Bergen, J.A. 1998. Cenomanian-Santonian calcareous nannofossil biostratigraphy of a transect of cores
drilled across the Western Interior Seaway. Stratigraphy andPaleoenvironments of the Cretaceous Western Interior
Seaway, USA. SEPM Concepts in Sedimentology andPaleontology, 6: 59-77.
Burnett, J.A. (with contributions from Gallagher, L.T. &Hampton, M.J.) 1998a. Upper Cretaceous. In: P.R. Bown
(Ed.). Calcareous Nannofossil Biostratigraphy. British
Micropalaeontological Society Publications Series.
Chapman & Hall/Kluwer Academic Publishers, London:132-199.
van Dongen, B.E., Talbot, H.M., Schouten, S., Pearson, P.N. &Pancost, R.D. 2006. Well preserved Palaeogene andCretaceous biomarkers from the Kilwa area, Tanzania.
Organic Geochemistry, 37: 539-557.
Kennedy, W.J., Gale, A.S., Bown, P.R., Caron, M., Davey, R.J.,
Gröcke, D. & Wray, D.S. 2000. Integrated stratigraphy
across the Aptian-Albian boundary in the Marnes Bleues, at
the Col de Pré-Guittard, Arnayon (Drôme), and at Tartonne(Alpes-de-Haute-Provence), France: a candidate Global
Boundary Stratotype Section and Boundary Point for thebase of the Albian Stage. Cretaceous Research, 21: 591-720.
Lees, J.A. 2002. Calcareous nannofossil biogeography illustrates palaeoclimate change in the Late Cretaceous Indian
Ocean. Cretaceous Research, 23: 537-634.
Lees, J.A. & Bown, P.R. 2005. Upper Cretaceous calcareous
nannofossil biostratigraphy, ODP Leg 198 (Shatsky Rise,
northwest Pacific Ocean). In: T.J. Bralower, I. Premoli Silva& M.J. Malone (Eds). Proc. ODP, Sci. Results, 198: 1-60
(online). http://www-odp.tamu.edu/publications/198_SR/
VOLUME/CHAPTERS/114.PDF
Nicholas, C.J., Pearson, P.N., Bown, P.R., Dunkley Jones, T.,
Huber, B.T., Karega, A., Lees, J.A., McMillan, I.K.,
O’Halloran, A., Singano, J.M. & Wade, B.S. 2006.
Stratigraphy and sedimentology of the Upper Cretaceous to
Paleogene Kilwa Group, southern coastal Tanzania. Journalof African Earth Sciences, 45: 431-466.
Pearson, P.N., Ditchfield, P.W., Singano, J., Harcourt-Brown,
K.G., Nicholas, C.J., Olsson, R.K., Shackleton, N.J. & Hall,
M.A. 2001. Warm tropical sea surface temperatures in the
Late Cretaceous and Eocene epochs. Nature, 414: 470.
Pearson, P.N., Nicholas, C.J., Singano, J.M., Bown, P.R., Coxall,
H.K., van Dongen, B.E., Huber, B.T., Karega, A., Lees, J.A.,
McLeod, K., McMillan, I.K., Pancost, R.D., Pearson, M.,
Msaky, E. 2006. Further Paleogene and Cretaceous sedimentcores from the Kilwa area of coastal Tanzania: Tanzania
Drilling Project Sites 6-10. Journal of African Earth
Sciences, 45: 279-317.
Pearson, P.N., Nicholas, C.J., Singano, J.M., Bown, P.R., Coxall,
H.K., van Dongen, B.E., Huber, B.T., Karega, A., Lees, J.A.,
Msaky, E., Pancost, R.D., Pearson, M., Roberts, A.P. 2004.
Paleogene and Cretaceous sediment cores from the Kilwa
and Lindi areas of coastal Tanzania: Tanzania Drilling
Project Sites 1-5. Journal of African Earth Sciences, 39: 2562.
Self-Trail, J.M. 1999. Some new and rarely documented Late
Cretaceous calcareous nannofossils from subsurface sediments in South Carolina. J. Paleont., 73: 952-963.
Young, J.R., Bergen, J.A., Bown, P.R., Burnett, J.A., Fiorentino,A., Jordan, R.W., Kleijne, A., van Niel, B.E., Romein, A.J.T.
& von Salis, K. 1997. Guidelines for coccolith and calcareous nannofossil terminology. Palaeontology, 40: 875-912.

New Late Cretaceous calcareous nannofossils...Tanzania

Plate 1
Calciosoleniaceae, Stephanolithiaceae

53

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Plate 2
Microrhabdulaceae, Braarudosphaeraceae, incertae sedis

New Late Cretaceous calcareous nannofossils...Tanzania

Plate 3
Incertae sedis, placoliths (uncertain affinity), Kamptneriaceae, Rhagodiscaceae

55

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Plate 4
Rhagodiscaceae, incertae sedis, Stephanolithiaceae, Axopodorhabdaceae

New Late Cretaceous calcareous nannofossils...Tanzania

Plate 5
Biscutaceae, Cretarhabdaceae, Prediscosphaeraceae

57

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Lees

Plate 6
Eiffellithaceae, Chiastozygaceae, Kamptneriaceae

New Late Cretaceous calcareous nannofossils...Tanzania

Plate 7
Chiastozygaceae

59

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Plate 8Chiastozygaceae

New Late Cretaceous calcareous nannofossils...Tanzania

Plate 9Chiastozygaceae, Calyptrosphaeraceae

61

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Lees

Plate 10
Calyptrosphaeraceae

New Late Cretaceous calcareous nannofossils...Tanzania

Plate 11
Calyptrosphaeraceae

63

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Lees

Plate 12
Calyptrosphaeraceae

New Late Cretaceous calcareous nannofossils...Tanzania

Plate 13
Calyptrosphaeraceae

65